Arctoa (2002) 11: 63-80 THE SAXICOLOUS MOSS FLORA OF THE MIDDLE URALS ÔËÎÐÀ ÌÕΠÑÊÀËÜÍÛÕ ÎÁÍÀÆÅÍÈÉ ÑÐÅÄÍÅÃÎ ÓÐÀËÀ IRINA L. GOLDBERG1 È. Ë. ÃÎËÜÄÁÅÐÃ1 Summary The saxicolous moss flora of the Middle Urals, the lowest part of the Ural Mts extending from 55054’ N to 59015’ N, is studied. It is found to be quite specific, comprising 110 species inhabiting exclusively or predominantly rock outcrops. Number of their localities in the Middle Urals, types of bedrock, altitudinal, regional and general distribution are analyzed. Annotated list of moss species with comments on morphology and anatomy of some taxa is provided. Ðåçþìå Èçó÷åíà ôëîðà ìõîâ ñêàëüíûõ îáíàæåíèé Ñðåäíåãî Óðàëà – ñàìîé íèçêîé ÷àñòè Óðàëüñêèõ ãîð, ïðîñòèðàþùåéñÿ îò 55054’ äî 59015’ ñ.ø. Îíà ÿâëÿåòñÿ äîâîëüíî ñïåöèôè÷íîé, âêëþ÷àþùåé 110 âèäîâ, ïðîèçðàñòàþùèõ èñêëþ÷èòåëüíî èëè ïðåèìó- ùåñòâåííî íà ñêàëüíî-êàìåíèñòûõ ñóáñòðàòàõ. Ïðîàíàëèçèðîâàíà ÷àñòîòà èõ âñòðå÷àåìîñòè, ðàñïðåäåëåíèå ïî âûñîòíûì ïîÿñàì è ãîðíûì ïîðîäàì íà Ñðåäíåì Óðàëå, ðåãèîíàëüíîå è îáùåå ðàñïðîñòðàíåíèå. Ïðèâåäåí àííîòèðîâàííûé ñïèñîê âèäîâ; äëÿ íåêîòîðûõ ðàññìîòðåíû àíàòîìî-ìîðôîëîãè÷åñêèå îñîáåííîñòè. Rocks on top of hills and mountains, cliffs The climate is temperate, more continental along river banks are widespread in mountainous on the eastern slope. The coldest month is Jan- regions. Rock outcrops are characterized by ex- uary with a mean air temperature of –16-170C, tremely fragmented and variable microenviron- while the warmest month is July with a mean ment providing for the highest diversity of habi- of +16-170C. The annual amount of precipita- tats for bryophytes. Besides, rock outcrops are known tion is 400-500 mm (Kuvshinova, 1968). to be places where lots of rare, endemic and relic Most of the territory is covered with boreal plant species occur. The flora of vascular plants forests. Dark coniferous woods of Picea obovata growing on various rock outcrops in the Middle and Abies sibirica predominate on the western Urals, and their geographic distribution have been slope, while pine (Pinus sylvestris) forests pre- well studied, while the knowledge of saxicolous vail on the eastern slope. The very south of the mosses of the area have remained very poor. Middle Urals is covered with forest-steppe plant The Ural Mountains is a long range located communities (Krasnoufimsky district) and birch- along the boundary between Europe and Asia. aspen (Betula verrucosa, Populus tremula) for- It is divided into five parts: the Polar, Sub- ests (Kamensky district). Areas above 650-700 Polar, Northern, Middle and Southern. The Mid- m a.s.l. belong to the subalpine belt with fir and dle Urals extends from 55054’ N to 59015’ N spruce open stands, sometimes with birch (Betu- (Borisevich, 1968) (Fig. 1). It is the lowest part la tortuosa), meadows and rock fields. Tundra of the Ural Mts., with its western slope on the vegetation is formed on the most windy slopes territory of Perm Province, and the eastern one (Igoshina, 1952; Ignatova et al., 1995). in Sverdlovsk Province. The relief is rather gentle, Wide spectrum of rock types is presented with average heights of 300-600 m a.s.l. The in the Middle Urals. According to the concen- highest point is Middle Baseg Mt. (994.8 m) – tration of SiO2 these are divided into acidic the main peak of the Basegi range situated in (65-80% of SiO2 – granite, quartzite), medium the north-western part of the area. (52-65% of SiO2 – diorite, andesite), basic (about 1 – Institute of Plant and Animal Ecology, 8. March St. 202, Yekaterinburg 620144, Russia – Èíñòèòóò ýêîëîãèè ðàñòåíèé è æèâîòíûõ ÓðÎ ÐÀÍ, óë. 8 Ìàðòà, 202, Åêàòåðèíáóðã, 620144, Ðîññèÿ 64 IRINA L. GOLDBERG 50% of SiO2 – basalt, diabase, gabbro, pyroxen- Campylium hispidulum, Schistostega pennata. ite, chlorite and talc schists), and ultrabasic (less 11 moss species (Cynodontium strumifer- than 45% of SiO2 – dunite, serpentine). They um, Dichodontium pellucidum, Dicranum spa- are all characterized by rather low concentra- diceum, Homalia trichomanoides, Hygrohypnum tions of Ca. Most of these rocks (granite, andes- ochraceum, Oxystegus tenuirostris, Plagiotheci- ite, diorite, basalt, diabase, gabbro, pyroxenite, um cavifolium, Platydictya jungermannioides, dunite) are volcanic, and some (quartzite, ser- Platygyrium repens, Pohlia cruda, Racomitri- pentine, schists) are metamorphic. Sedimentary um canescens) have been found predominantly rocks are also common – these are limestones on rock outcrops, with a few collections on oth- consisting mostly of CaCO3. er substrates. 99 species and 5 varieties belong- In 1996-2000, I collected saxicolous mosses ing to 62 genera of mosses are known to grow in Sverdlovsk Province: on the granite outcrops exclusively on rock substrates in the Middle close to Yekaterinburg, limestone cliffs in the Urals. These are obligate saxicolous species, and valleys of the Ufa, Serga, Chusovaya, Iset, Rezh I would like to underline that they grow not rivers and their tributaries, basic and ultraba- only on bare rock surfaces (as one could think), sic rock outcrops along the Sysert and Iset riv- but also on humus and soil accumulating in ers, and on tops of a few hills. The specimens crevices and niches, and on cliff ledges. are placed in the Herbarium of Institute of Plant The rest of this article will be devoted to the and Animal Ecology (SVER) in Yekaterinburg. diversity and distribution patterns of moss spe- I also examined other collections of mosses from cies found in the Middle Urals exclusively or the Middle Urals housed there and in the her- predominantly on rock substrates. baria of the Ural Teachers’ Training State Uni- Number of localities in the Middle Urals. versity in Yekaterinburg (which still has no For each species I counted the number of sites official acronym and called below as UTTSU), where it was recorded. Species found at 1-10 Perm State University (PERM), Moscow State localities were regarded as rare, at 11-20 local- University (MW), Main Botanical Garden ities – common, at more than 20 localities – (MHA) in Moscow, and of the Botanical Insti- frequent. Only 16 species of saxicolous mosses tute (LE) in St. Petersburg. Additional data were (Anomodon viticulosus, Cynodontium strumifer- obtained from Krylov (1885), Navashin (1888), um, Distichium capillaceum, Hedwigia ciliata Syuzev (1899, 1909), Gorchakovsky & Arkhipo- and others) are frequent, and 9 (Anomodon va (1964), Sitnikova (1976), Dyachenko (1987, longifolius, Cynodontium tenellum, Grimmia 1990, 1996, 1997a, b), Trotsenko (1990), Ignatov affinis, Encalypta rhaptocarpa, Hypnum vaucheri, & Maslovsky (1991), Ignatova et al. (1995), Pseudoleskeella tectorum, Taxiphyllum wis- Dyachenko et al. (1996), and Bezgodov (2002). grillii, Tortella tortuosa) are common on rock The saxicolous moss flora is known from 85 outcrops in the Middle Urals; that makes a bit localities in the Middle Urals (Fig. 1, Appendix 1), more than 20 percent of the flora. All of them most of them are situated in the central part of are known from both Perm and Sverdlovsk the eastern slope. The saxicolous flora comprises Provinces. Other taxa (85) are rare. Of them, 16 190 species and 5 varieties. 80 taxa facultatively species have been found only in Sverdlovsk grow on rock substrates (Table 2), and I am not Province, and 26 taxa only in Perm Province. A sure that the list is complete. Most of these species very high percent of rare species in the saxi- are widespread in boreal forests, where they in- colous moss flora of the Middle Urals can be habit soil and decaying wood or grow as epi- explained by their ecological features, such as phytes. Some are typical of disturbed habitats: altitudinal distribution in the Middle Urals and open soil in woods and fields, sides of roads and types of rock they inhabit, as well as geograph- ditches, pits under roots of fallen trees. Hygro- ic distribution – in the Ural Mts. and Eurasia. phytes from wet habitats (flood-plains, bogs, mea- Altitudinal distribution. Although highlands dows, and boggy forests) can inhabit boulders in do not cover vast areas in the Middle Urals, there river valleys. Some facultative saxicolous taxa are are 9 rare taxa (Andreaea rupestris var. papillosa, rare in the Middle Urals: Amblystegium varium, Anomodon rugelii, Dicranowesia crispula, Grim- Brachythecium erythrorrhizon, Buxbaumia aphylla, mia elatior, Hygrohypnum alpestre, Iwatsukiella The saxicolous moss flora of the Middle Urals 65 Fig. 1. The study area. leucotricha, Lescuraea saxicola, Pseudoleskea radi- cosa, Racomitrium sudeticum) reported only from the subalpine belt. 35 species (Amphidium lap- muehlenbeckii). In general, moss species diversity ponicum, Bartramia ithyphylla, B. pomiformis, is the highest on limestone, and the lowest on gran- Cynodontium strumiferum, C. tenellum, Grim- ite. mia muehlenbeckii, Paraleucobryum longifolium, Type of bedrock the species is restricted to Pohlia cruda and others) are known from both can partly explain its altitudinal distribution. For forest and subalpine belts. Among them, Andre- example, there are no limestone outcrops above the aea rupestris var. rupestris, Racomitrium micro- timberline in the Middle Urals, and all calcicole carpon and Polytrichastrum alpinum are very moss taxa are known only from the forest belt. common in the subalpine belt and rare at low Distribution in the Ural Mountains have elevations. Only one collection of Anomodon been analyzed using the results of my own in- attenuatus has been made above the timberline, vestigations in the Northern (Goldberg,